Wednesday, January 11, 2017

This post might piss off a few people… including some regular readers of the blog. But if you don't know it already you shouldn't come here expecting not to be challenged in some ways!! And sometimes we all - including myself - need the piss taken out of us a bit.

The battle of out times - scales versus feathers. I imagine some epic Lord of the Rings battle in middle earth: scale loyalist Orcs massing against feather nazi elves (feather nazi elves lolz). Various banners of the opposing forces drawn up with resounding battle cries and triumphant bellows of warfare. I paint this picture to obviously poke fun at members of both parties and to set a tone of jovial banter. Cuz it is kinda funny when you step back from it and realize 99.999% of people don't really care... Of course I care and so do my readers but I think discussions would be more fruitful if both factions stood down a bit and realized that there are resounding points that both sides make and, as I will highlight in this piece, there are some points and arguments to be made that have largely been circumscribed in the exchange.But if you find yourself getting heated while reading this just pinch yourself, and then remind yourself that in a world of of growing nationalism, fundamentalism, misinformation campaigns, cyber-attacks, and frank talks of nuclear bombardment what dinosaurs wore for apparel does not really matter much...

The long running scale versus feather argument, largely played out online and largely converging on theropods (especially T. rex) integument has become mired down into two factions.

The pro-scale minions, seemingly on the defeat and defensive suffering wave after wave of feathered theropod discoveries and to top it off ornithischians with filaments. Most damning of all, Yutyrannus a largish tyrannosauroid with abundant feather impressions over large parts of the body. This and other feathered basal tyrannosauroids creating the valid and likely position that more derived tyrannosaurids also sported (some amount of) plumage. Still the pro-scale contingent has reason to remain vocal. Scale or scutes is all we really have for many lineages of dinosaurs; Carnotaurus; derived tyrannosaurids and even T. rex. Scales and scutes certainly have a big role to play in theropod and dinosaur integument and they did so for the duration of the Mesozoic.

Diametrically opposed to the scale loyalists are the self proclaimed (I kid you not there is an actual satirical deviantart group) ... "feather nazis". Theirs is the grand and exciting enfluffening movement where everything from sauropods to ceratopsids is gowned and strutted with honest to goodness flowing manes of insulatory integument. Feathers for all!! Feathers right down to the base of dinosauria!! What was considered overly speculative by many in the 90's has now metastasized into a dogma all of its own. What the feather nazis lack in outright evidence they more than make up for in passion and online advocacy (shaming?).

A more succinct analysis would be that nothing triggers a scale loyalist like a fluffy T. rex and nothing triggers a feather nazi like a JP stylized raptor.

My main beef with both factions is that they have perpetrated a bit of a false dichotomy. Lost in the garbled rhetoric of both parties is a third option that has largely been eschewed, silenced, or simply ignored. That third option is nekkid skin itself. As I will lay out nekkid skin perhaps highly keratinized or fleshy - the most basal, germinal layer of all - on the head and even neck in many cases is a quite defensible option for theropods and perhaps even other dinosaurs. But first I must slaughter some feather nazis.

Now my issue with the "feather nazis" is not one of complete disagreement. I think there is good reason to argue feathers or at least stage one quilly type things all the way back to the origin of theropods or more basally to stem archosaurs. However somewhat embedded in this suggestion is that such insulatory "protofeathers", "dino-fuzz" or "quilly" type structures are wholly and completely superior to "naked" scutes and scales in terms of insulatory potential. Or just negating the potential for thermoregulatory capacity in scaley structures at all and dismissing them as "inferior" or harkening back to more "primitive" gestalts.

I believe that scale loyalists have really missed or conceded an important argument in favor of abundant and diverse scaley type integument in dinosaurs. S.I.G.I.L. "subdermal interstitial gridded insulatory layer" an idea I suggested here a while back makes the case that scales, scutes, even osteoderms, have important potential for insulatory adaptions. Here I suggested that small pockets underneath the outer layer of skin in naked hided dinosaurs created an insulatory vacuum sealed layer when blood was withdrawn internally, basically the same concept of a thermos or people insulating their house with bubble - wrap. So I am not so sold that naked hided dinosaurs are a bad thing of lacking in insulator capacity even at small sizes or in cold climates. If certain basal prosauropods sported insulatory coats (no evidence yet) or basal ornithischians that we know did (Kulindadromeus yes!!) S.I.G.I.L. might offer a reason why this coat was lost and basically not needed in more derived forms like hadrosaurs, ceratopsids, ankylosaurids etc. etc. or maybe never there to begin with!!

No, my main beef with the "feather nazis" is not just a certain arrogance and condescension in their ranks but a certain amount of committing "sins of omission". We should always be careful to consider are we making the best argument or are we asserting what we want. And I don't really care if you are asserting what you want but don't pretend not to be!!

Time to take the feather nazis down a peg or two.

For best effect the arguments of the feather nazis should be read in the voice of Gollum. Like Gollum, whose mind was poisoned by the ring and all that it brought, the feather nazis have also came in possession of an ancient and powerful artifact - the feather - that has likewise poisoned and twisted their mind.

"But, my precious, feathers protect from harmful UV rays..."
This is true. But it is a bit of a half truth. What it is suggesting is that if it was not for feathers naked skin would be burnt to a crisp like extra crispy Kentucky Fried Chicken. This is simply not true and you know what protects from UV the best... melanin found in feathers and skin. You don't really need feathers to shield from UV rays, simply put melanin in your skin (as nekkid skin birds do already) and the problem is solved.

"Yes but feather have unique insulatory properties that keep birds thermally protected from extremes in heat or cold my precious..."

This is true. But again the assumption is that scales or scutes (esp. as I outlined in my S.I.G.I.L. hypothesis) could not do that job aptly too. Scales/scutes are rather unexplored as unique thermoregulatory organs in their own right and need not be assumed as vastly inferior to feathers in terms of insulatory properties in light of the lack of study on them. Especially if we assume a lower middle metabolic rate, scute and scale having dinosaurs might have done just fine… sort of like armadillos (more on xenarthans later).

A BIG problem in feathers covering the totality of large archosaur bodies in hot climates (read large dinosaurs in hot climates) is the issue of heat exchange. If the whole or most of the body is draped in feathers - which have excellent insulatory capacity - and body heat rises too high there is nowhere to shed this excess heat. Except, of course, by allowing for large tracts of scaley or nekkid skin (as modern birds do).

"Precious, you have forgot about the heat shedding abilities of the avian lung system..."

Ah... the coup argument against large tracts of naked skin or scales for large feathered dinosaurs in hot climates. The problem is that the poultry ranchers of California don't agree with you.

A bit ago I taught traffic school and one of the more obscure laws on the books is one concerning objects that can fall out of a moving vehicle and not be considered litter (absolving one of a $1000 fine). Clear water and feathers from live birds.Spilling Loads and Damage to Highway.(VC §§23114 and 23115) It is against the lawto operate on the highway a vehicle which is improperly covered, constructed, or loaded so that any part of its contents or load spills, drops, leaks, blows, sifts, or in any other way escapesfrom the vehicle. (Exception: clear water or feathers from live birds.)

Turns out that transporting live poultry requires open air containers in order for adequate airflow and avoiding heat stress. In fact a cursory investigation into the issue of heat stress and birds (just google birds overheat) is rife with accounts of heat induced stress and death.The mighty avian lung system might not be the panacea we assumed it was in terms of preventing overheating. Additionally that turkeys and chicken have both avian lungs and fleshy display structures shows that even with both of these adaptations overheating still occurs. Overheating occurs quite a lot in birds actually. So I would not put much merit in this argument to counter abundant and extensive tracts of nekkid skin or scales.

The better bet for being a large feathered archosaur in hot climates is to clear up large tracts of nekkid skin on the face, around the eyes, the neck, even the chest. Get nekkid. Like Lappet-faced vultures. Or marabou storks. Or ground hornbills.

Pelage Loss: For Some But Not All?

There is one important concession that I have never seen feather nazis make. If insulatory apparel goes way back to the base of dinosauria or even earlier then we must presume that it was secondarily lost in some lineages : sauropods, hadrosaurids, ceratopsids are the most commonly cited potential examples of this. However if we presume that they could have lost their plumage then why can't we do the same for large tyrannosaurids? I don't think it is an argument that the pro-scale loyalists have been vocal enough about. I don't myself necessarily agree with it but I also suspect that tyrannosaurids had a heck of a lot of scale coating and nekkid skin on them.

In fact if people are allowed to have the boldness and audacity to depict "feathered" sauropods (and I believe that they should although I don't necessarily think it likely) they should also be allowed the boldness and audacity to depict a giant, perhaps mainly scaled and nekkid skinned maniraptoran that has lost almost all of its plumage. Developmentally it could happen. Personally I doubt it and I think a wooly sauropod and a mainly featherless maniraptoran are long shots, but if we allow speculation to one extreme why not the other? Boy would that trigger the feather nazis.

The Nekkid Skinned Archosaur

The argument is rather simple and straight forward. More so than that it is bolstered by the patterns we see in living feathered archosaurs. Go read this excellent online summary from an ornithology class on avian integument

Ye shall know it when ye sees it...

What was once completely scaled shall henceforth be completely feathered...From the tip of the tail to the tip of the noggin' ye shall know only feathers until...Conditions henceforth arise that necessitate a shedding of select feathers revealing...a nekkid primordial skin... rippling, oozing, and stimulating with a lustful passion for display!!

Or to put it another way:

1) Scaled theropod evolves feathers. Not just a few feathers, but a full on coat covering the whole body as evo-devo studies suggest. Because feathers and scales are somewhat competitive developmentally what was skin covered with scales now just becomes skin covered with feathers.

2) Later on certain members of this feathered tribe get big or move to hotter climes or both. In order to shed heat, feathers retract from certain areas - the head, feet, the tail. In certain spots that are prone to impact or abrasion against the ground scales reemerge. This is exactly like the "scaled" feet and legs of modern birds. In other areas the feather retract to reveal nekkid skin.

3) And when modern archosaurs shed the feathers on their head, around eyes, the neck... when birds get nekkid they freaking play with that skin in all kinds of weird and wondrous ways. Big time freaks.

4) Bird skin is thin. If you eat some bbq chicken it comes with the skin. If you order some bbq beef it does not come with the skin. So that when feathers are lost one potential way to toughen up skin is to add more layers, wrinkles and weird growth type thingies. Cancerous looking caruncles and tufts of skin, waddles.

helmeted Guinnea fowl. credit Bob CC3.0

Abyssinian ground hornbill. female. credit Jerry Thompson CC2.0

credit Eric Kilby wattled crane CC2.0

A frequent dismissal of this generous endowment of soft tissue display/thermoregulatory structures onto non-avian theropods is parroted at me as such:

"Such skin display structures are not that widespread in birds, in fact statistically speaking we probably have not even found a dinosaur with such display structures."

For starters it is a text book case of bad statistics. While it is true that if you measured the prevalence of naked skin/fleshy display structures in modern birds you would find a fairly low percentage - I would hazard less than 5% of birds if not less have such structures - but numbers without analysis and context is meaningless. What you have to do is look at the likely ecology of dinosaurs and find what birds best match. And if you look at the birds that best match dinosaurs in ecology I would pick the ones that are large (for a bird that would be partridge size), mainly terrestrial or at least feed on terrestrial items, and live in temperate to hot climates. I would not put aquatic birds and passerines high up in ecological correspondence but even some passerines and waterfowl sport naked skin structures. Among the birds I would put forth closest to non-avian theropods ratites, galliformes, "vultures", storks, hornbills, cranes and especially ground hornbills as some of the best matches. Once you start looking at the birds that best fit non-avian theropod dinosaurs ecologically and environmentally you start to see that such nekkid, fleshy, and skin derived display/thermoregulatory features are quite common. Nothing is 100% and not all of these birds fit the bill - but a large enough proportion of them do that we should be highly confident that such displays were at least as common in feathered dinosaurs. More common I would argue because non-flighted dinosaurs could afford heavier skin structures while birds have to be a bit more thrifty in terms of such heavy building material. Also, the compacted lifespan of dinosaurs and hyper-competitive sexual politics of dinosaurs would encourage the elaboration of skin derived display structures on both sexes. Also, and most importantly, because they tend to piss off some feather nazis and anything to take the piss out of them.

In fairness there are notable exceptions to this large, terrestrial bird with naked skin/fleshy faced pattern. Biology is after all notoriously squishy and as soon as you push down on something and proclaim "this is the answer" exceptions start to arise. Kori bustards are fully feathered and siriemas only have a little eye patch of naked skin. Cranes don't have loads of fleshy skin derived facial structures but they are highly migratory (remember skin weighs more than feather) and principally temperate. Far be it to me to insist on fleshy faced, skin display structures on all theropods even maniraptorans - I don't really see that this issue is as clear cut as even I would like it to be. Herons are another group that doesn't fit the trends as they have fully feathered necks and faces. However in their case they need to use rapid neck strikes to catch prey and skin weighs more than feathers so better to keep the feathers for display. Certainly smallish microraptorines types likely had fully feathered faces, or species that lived in cool temperate climes (i.e. Liaoning). But gigantic oviraptorines, dromies, ornithomimids, tyrant lizards, therizinosaurs, megaraptors and other largish coulurosaurs I would really hedge my bets towards such gnarly skin & flesh derived facial and forequarter display/thermoregulatory structures as fairly common. How far back you want to take such skin derived structures depends somewhat on how far back you want to take feathers in theropods. In other words the evolution of feathers begets naked skin derived structures.

The other point lost entirely on some is that I don't have to speculate about nekkid skin and derived structures - we already have proof.

Pelecanimimus polyodon. How come nobody talks about Pelecanimimus anymore? This is a basal ornithomimosaur and therefore a coelurosaur and therefore feathered. The soft tissue preserved around the head and neck as interpreted by the authors is very telling. No scales present. No feathers present. Instead impressions revealing wrinkles (possibly desiccated caruncles?) of skin lacking integumentary structures i.e. no feathers or scales. A soft tissue crest or flange of skin or keratin on the back of the head. A "pelican like" gular pouch. Just as I would predict and just as we should expect for a good number of feathered dinosaurs in hot climates. Soft tissue structures and nekkid skin. Also Edmontosaurus.

*update Darren Naish (see comment) let me know Pelecanimimus soft tissue interpretations are a bit of a wash. What did I say about taking the piss out everyone (including myself)? Maybe the wiki page needs an update too...

Of course if an ornithomimid the size of Pelecanimimus is showing a tendency to clear up feathers for nekkid skin and display structures you can imagine how far Deinocheirus could have taken things...

Let's not to forget to feed the rumor mill. If you dig deep enough there are some accounts of nekkid facial skin on tyrannosaurids and ol' sexy rexy himself. Now I don't want ol' Rexy to dominate this discussion but there are consistent rumors of "plucked chicken" looking skin, bits of skin with reduced scalation and completely naked skin. Feather nazis should rejoice at the notion of naked skin because as I highlighted earlier naked skin implies it was once feathered and there is likely reduced feathering somewhere on the body. But the sheer amount of retrieved "gila-monster" like scalation does suggest that scales were an important and potentially widespread ectodermal covering for tyrannosaurids. Probably more widespread than in the secondarily evolved scalation of modern birds.

I and a few other artists and thinkers that are increasingly embracing nekkid skin on the head & forequarters of theropods are a threat to both the scale loyalists and some feather nazis. I put scale loyalists in a frenzy because this look completely alters the iconic typography of how a theropod head "should" look - primarily portrayed as a tightly adhering scaled shellac over the theropod skull. And I know I really get under the skin of feather nazis because I argue - based on what modern birds do - that the attractive, graceful, and highly feathered facial countenance as depicted in many non-avian theropods is substantially different than what they have become beholden to.

Some feather nazis deserve to be put in their place a bit because some are guilty of the same crimes that they charge the scale loyalists with. What am I saying? Feather nazis often charge the scale loyalists with being emotionally and culturally clinging to an old outdated 90's nostalgia - the T. rex and raptor of Jurassic Park fame. However from my point of view when I advocate abundant fleshy and skin display features in theropods - inspired by birds of course - who do I get the most push back from? Feather nazis. Why? Even though nekkid skin presupposes a fully feathered ancestral state once you go the path of nekkid skin then you pretty much have to go the path of outrageous skin derived structures. And this completely ruins the fully feathered and attractive plumage depicted on the head of most feathered dinosaurs and calls into question how ubiquitous the "ground hawk" look was for dromies. Feather nazis don't like me for primarily the same reason scale loyalists don't like feather nazis. I call into question a certain look that they have grown fond of. Feather nazis can't outright dismiss me based on an adherence to past interpretations as they can do with the scale loyalists.

Confounding the situation is that many paleoartists give theropods a veneer of feathery type insulation but capitulate to the scale loyalists by giving theropods a scaly head. Why do they do this? If you look at what modern birds do when they lose the feathers on the head all you are left with is naked, not scaly, skin.

So far I have taken it a bit easy on the scale loyalists. That is gonna change.

(best read in an awesome - bro voice, whatever that means to you)

"But, yeah, hey man what about like Carnotaurus it had like scales like top to bottom man. Fully bruh."

Ahh I'm glad we get to talk about Carnotaurus. And it was literally shellacked with scales, scutes, nodules galore. Not a feather to be found. Carnotaurus is indeed the clarion call for scale loyalists and they should not budge from what it tells us - just as feather nazis should not budge from what Yutyrannus tell us. What I really want to talk about is the face of Carnotaurus because, let's be real, that's what we really care about right? The question I ask is did Carnotaurus - and by extension other abelisaurids - actually have a scaly face?

Yeah I am actually serious when I say this. Carnotaurus may have not had a scaly face, it may have indeed had a croc face.

Check out this nice summary piece, croccrack, showing that the assumed scales on the heads of crocs are actually cracked skin that forms embryologically from the underlying bone pressing against it.

Now, I am not one to lean to fully on the extant phylogenetic bracket to infer soft tissues, and I won't start now, but one of the interesting observations is that no living archosaurs have scaly faces. Birds don't although the wood stork comes close with a highly keratinized face. And crocs actually do not have scaly faces, they have a very tight fitting skin that cracks and creases along defined planes of stress embryologically creating the illusion of very irregular scales. Yep, that was something that really threw me for a loop when I first came across that info on Jaime Headden's blog. But it is true, crocs have scales everywhere else on the body but not on the head. Could this have something to do with better use of ISO's on the snout and head... possibly. Stay posted on that thought.

CC3.0 credit A Stranger in the Alps

It is one of those frustrating aspects of paleontology that the holotype of Carnotaurus supposedly contained remnants of the facial skin but they were prepared away... gripes!!

"Originally, the right side of the skull also was covered with large patches of skin - this was not recognized when the skull was prepared and these patches were accidentally destroyed." wiki citing Czerkas 1997

However there are at least some descriptions available of what the facial skin of Carnotaurus looked like which included bits from the lower left jaw and right side of the head:

"Scalation was similar across different body parts with the exception of the head, which apparently showed a different, irregular pattern of scales." wiki citing Donald F. Glut Carnotosaurus: Dinosaurs 3rd edition

Our first clue is that the "scales" of the head were described as "different" from the rest of the body. Hmmm why would that occur? Does the scalation on a monitor lizard head differ substantially from the rest of the body scales? Our second clue is that the scales of the head were described as "irregular"... where have we heard that descriptor before? Oh yeah, the "scales" on the heads of crocs (actually cracks in scaleless skin) are also described as "irregular". Our third clue is that the skull of both crocodiles and abelisaurids are highly rugose, bumpy, and just gnarly looking. The tight fitting skin that covered the skull was equally keratinized, rugose and just gnarly looking. Our fourth clue has to do with why - adaptationally - would crocodiles lose scales on the head? It might have something to do with allowing better transmission of data to those integumentary sensory organs on the head of crocs. Forgive me for lacking a reference but I thing I came across the suggestion of that idea on Mark Witton's blog once... need further info. Regardless I think it worth further investigation.

Remember when it was common sense that crocs had scaly heads? Why, if we for so long assumed that, is it not a definite possibility that the first impulse when confronted with the preserved facial integument of Carnotaurus could the researchers have not made a similar mistake?

Fucking rad, right? Much props to J.W. Kirby for this Aucasaurus with a keratinized but non-scaly head

J.W. Kirby was nice enough to lend his talents for this depiction of an Aucasaurus with a keratinized mug. I think it works exceptionally well and does a good job of splitting the difference between a wood stork and a croc. Visit his deviantart: KirbyniferousRegret

Am I suggesting that abelisaurids lost the scales on their face to enact ISO's? Nope, although the idea of subaquatic abelisaurids has been toyed with, the swarm of data points towards terrestrial. So I don't think that they had ISO's but I do think that they had especially tactile faces. They had tough faces for sure - highly keratinized - but they also had tactile and sensitive regions, especially along the oral margin i.e. the lips. And I would suggest that they had such tactile lips to better react and position themselves with regard to prolonged and dangerous engagements with prey. Their teeth, their jaws their whole predatory apparatus was set up to bite onto and hold onto prey - not slice through it like the vast majority of other theropods. Dispensing with scales on the head allowed for a more tactile and also tough and keratinized predatorial and cranial adaptation to flourish. And also face biting.

Wood Stork. credit Sandy Sharkey/Great Backyard Bird Count

So if you want to draw abelisaur heads a nice informed speculative venture would be to look at crocs and wood storks for inspiration and add some tough but tactile lips and thickened gums!! I am actually converging on a perception that the upper lips of abelisaurids were very keratinized and did not hang low - almost on the path to lipless crocs. I think, however that the lower lips did hang laterally and inferior with a nice big keratinized gummy mess leading up to the teeth. Go bold and add some flesh antlers and outgroths off the cranial crest as suggested by the vessels on the top of the skull of Rugops!!
While we are on the subject of cranial scale reduction, keratinized skin, croc heads, and rugose skulls it is also interesting to wonder if the rugose skulls of other theropods that lived a very facial - tactile existence such as spinosaurids, unenlagines, other theropods and even pachecephalosaurids and some ceratopsids - may have dispensed with scales and opted for keratinized facial halos. Lots of potential candidates there.

I have some other things to say about scales and the potential loss of scales on the heads of theropods but I'm gonna save than for upcoming posts. Just know that I am not done with you yet "lizard-lipped" theropods...

Scale Loyalists Should Not Simply Concede Kulindadromeus or Psittacosaurus
Another issue I take with scale loyalists is that they simply seem to concede animals like Kulindadromeus and to a lesser extent other quilly ornithischians and possibly Concavenator to the feather nazi hordes in their blitzkrieg conquest. Don't stand down scale loyalists these animals have important stories to tell about scales too!!

credit Tomopteryx CC4.0 Kulindadromeus

Kulindadromeus is the best example so I will talk about this little Siberian maverick the most. Somewhat lost in the excitement about this ornithischian with I think 3(?!) different types of filamentous coverings is that it also had a diversity of scaley type coverings. I think 2 or 3 types of scale coatings. That is an important point not to be conceded by the scale loyalists. Think about where these scales were located. On the tail, the hands, the feet. The areas most liable for temperature oscillations. Scales do have an important story to tell and it likely has a lot to do with thermoregulation and scales remained very important for the duration of dinosaur history.

The quilly Psittacosaurus was pretty small, lived in a cool climate, could have probably grown a coat of insulating filaments. Certainly other similarly sized maniraptoran herbivores and omnivores in the same environment grew a full coat of feathers. So why not Psittacosaurus? We can rule out preservational bias as an argument so often posited for why feathers are "not preserved" as the Xixian beds preserve feathers/filaments well. This is a dilemma that feather nazis simply ignore by pretty much not addressing it or talking around it. It was scaled and those scales may have provided good enough thermoregulatory benefit that a more extensive filamentous coat was not needed.

An important thought to keep in mind with rear gut fermenting dinosaurian herbivores is that they walk around with big compost bins in their gut. Like compost does, rear gut fermentators release heat. This constant slow burn of heat could have allowed for a more reduced and basal insulatory apparel - scales dominating and perhaps in some instances supplanting filamentous coatings. If we look at scales as basically solar panels such animals could have further supplanted their heating costs by stealing and hoarding energy from the sun and environment. Muscular activity, solar panel scales, rearward heat creating fermentation, blood flow and perhaps a rather low and thrifty basal metabolic furnace could have all combined to create an especially energy thrifty and efficient design. An extensive plumage not really called for. This exceptionally thrifty design would have also begat exceptional ability to dedicate food resources to growth as opposed to heating the furnace, in line with the noted size of many dinosaurian herbivores.

Filaments and feathers may not so much as replaced scales but augmented them in some instances, especially in herbivorous dinosaurs.Xenarthran Dinosaur: The Further Back You Go, the More Complex Things May Get
I want to end this piece by pointing out where I think we might be heading. We might be heading into a zone of less certainty than more certainty with regard to integumentary structures in not just theropods but dinosaurs as a whole as well as other beasties like various stem-mammals of the synapsid gestalt that have also been receiving interest as goes soft tissue reconstruction. Now I realize that some might not take much solace in this. But I think that we should really begin to expect the unexpected. Before I let you know why I suspect that the integumentary patterns we might infer based on extant relatives might be a little too conservative let me post this abstract to a rather practical and succinct paper.

Abstract

In zoology it is well known that birds are characterized by the presence of feathers, and mammals by hairs. Another common point of view is that avian scales are directly related to reptilian scales. As a skin embryologist, I have been fascinated by the problem of regionalization of skin appendages in amniotes throughout my scientific life. Here I have collected the arguments that result from classical experimental embryology, from the modern molecular biology era, and from the recent discovery of new fossils. These arguments shape my view that avian ectoderm is primarily programmed toward forming feathers, and mammalian ectoderm toward forming hairs. The other ectoderm derivatives – scales in birds, glands in mammals, or cornea in both classes – can become feathers or hairs through metaplastic process, and appear to have a negative regulatory mechanism over this basic program. How this program is altered remains, in most part, to be determined. However, it is clear that the regulation of the Wnt/beta-catenin pathway is a critical hub. The level of beta-catenin is crucial for feather and hair formation, as its down-regulation appears to be linked with the formation of avian scales in chick, and cutaneous glands in mice. Furthermore, its inhibition leads to the formation of nude skin and is required for that of corneal epithelium. Here I propose a new theory, to be further considered and tested when we have new information from genomic studies. With this theory, I suggest that the alpha-keratinized hairs from living synapsids may have evolved from the hypothetical glandular integument of the first amniotes, which may have presented similarities with common day terrestrial amphibians. Concerning feathers, they may have evolved independently of squamate scales, each originating from the hypothetical roughened beta-keratinized integument of the first sauropsids. The avian overlapping scales, which cover the feet in some bird species, may have developed later in evolution, being secondarily derived from feathers.

Now, in several points in this piece I have referred to nekkid skin as the basal or germinal layer - that when scales or fur or feathers are lost what remains is pure skin. This study confirms that "inhibition (of the Wnt/beta-caratin pathway) leads to the formation of nude skin and is required for that of corneal epithelium." Which is pretty much what I have been suggesting happened quite a lot in dinosaurs that were formerly completely feathered but then - for a multiplicity of reasons - lost such feather coatings on parts of the head and potentially forequarters.

A more interesting bit of info from this abstract is the notion that - read modern - mammals and birds have ectoderm primarily programmed to grow feathers and fur respectively. Again, I want to reiterate that this is how modern bird and mammal skin appears to be programmed. But let us not assume that more basal (read extinct) forms were as hemmed in developmentally to form such structures. I first came across this notion in the above Jura post on the topic (feathers for one and all?) and there Jura referred to as a basic rule of thumb in the comments section: the regulatory pathways, the developmental constraints may have been a lot more lax and prone to slippage, reversions, and wholesale jumbled set ups than what the more modern derived and somewhat set in stone representatives would suggest. A lot more play. A lot more complexity. And a lot less certainty for anyone trying to arbitrate what is accepted and reasonable in paleoartistic representations.

Pink Fairy Armadillo. credit cliff1066 C3.0

*Update 1/22/17. I too hastily made the following comparison and it erroneous. I still think it worthwhile to consider more basal members of dinosaur lineages may have had a diversity of integumentary features - as Kulindadromeus suggests - but as several commentators pointed out the analogy is wrong. I will leave it up as is because I think it important to document things I get right and things I get wrong.

Consider xenarthrans - the strange and alluring mammals that include the extinct glyptodonts and giant grounds sloths, as well as armadillos, sloths, anteaters, probably some other weird ones too I am forgetting. I don't like to use the term "living fossil" but I think it fair to call these mammals the least derived or most basal among extant mammals. The most basal mammals but also the most diverse in terms of integumentary structures. Think about the weird scutes of armadillos and glyptodonts, the coarse hairs of anteaters, the osteoderm studded skin of giant ground sloths. Now in light of what was discussed in the last paragraph where I made the point that the most derived members of clades are "locked into" defined developmental pathways for respective integumentary structures. For extant birds this is feathers for mammals it is fur. But if you go further back into more basal forms in each group it is very probable that there was a lot more developmental wiggle room, a lot more play, and a lot less capacity to fully predict what was going on in terms of how much, where, and what type of integumentary structures were going on. Consider monotremes, also very basal and also diverse in terms of integumentary types. Pangolins and aardvarks as well.

Now, this is an area of needed study but in light of the basal nature of these animals and the breadth of integumentary structures that they evince I doubt it merely a coincidence that we see this congruence. That pangolins, monotremes, and xenarthrans showcase more diversity in integumentary structures than more derived and numerous ungulates, carnivorans, and rodentia combined is something to think about...

This notion that using highly derived extant relatives to ascribe dogmatic rules to their extinct relatives should not sit too well with both feather nazis or scale loyalists. And you know I like it when both sides of the divide get befuddled. That creates fertile living and creative space to speculate to truly bizarre and jarring amalgamations of quills, feathers, scales, scutes, keratinized skin, wattles, caruncles, frills, dewlaps, and other structures.

In fact we don't have to speculate. Kulindadromeus spells it right out for us being fairly basal and hosting a diversity of scutes, overlapping scales, and various types of filaments. Later ornithischians may have largely dispensed with pelage and just as derived modern birds and mammals are locked into a pathway of fur or hair more derived ornithischians became more and more locked into a developmental pathway of scales. You simply don't see derived ungulates with lumps of osteoderms in their skin do you? And while the maybe - maybe not ulnar quill nodes of Concavenator get most of the attention for that fantastic beast less widely reported is the unexpected diversity and size of large scales preserved with the body. The Mesozoic may have been rife with complex amalgamations of scaly, scutey, quilly, nekkid skinned, and feathery beasties. Enough to confound dogmatists on both sides of the aisle.

Its not about "the enfluffening" or the return of the retrosaur it is about the advent of the weirdening and we have been in it for a while now.

In summary both scale loyalists and feather nazis bring important points to the table but their rigidity sometimes veers into dogmatism.

Naked skin on many formerly fully feathered theropods is a thing, as evinced by Pelacinimimus. The reasons for clearing up naked skin on theropods (extinct and extant) is not fully understood, doubtless multifaceted, and as shown by species such as Kori Bustards and Siriemas not a 100% mandate or law. But it was a trend that was certainly common. The evolution of naked headed theropods depends in large part on how far back fully feathered dinosaurs goes. Whether or not tyrannosaurids sported naked skinned heads is up for grabs but a distinct possibility. Especially considering the tactile nature of their skulls, feathered pedigree, and several anecdotal records of scaleless or almost scaleless hide as remarked by several researchers (Phil Currie & Paul Sereno).

Because bird skin is so thin we should expect that when feathering is lost it should thicken up with wattles, caruncles, fleshy outgrowths, keratinized bits, and other diverse features we see in living naked faced birds. Expandable gular pouches, snoods, crests, and other features were also likely widespread. I would venture that these features are relatively more common in feathered non-avian dinosaurs than flight capable birds because 1) skin weighs more than feathers so flighted animals will tend to be a bit more thrifty with such features compared to grounded animals 2) the hyper competitive, hyper condensed mating politics of dinosaurs would stimulate outrageous, expensive, and elaborate features. Dinosaurs - unlike sea turtles, crocodilians, and long lived monogamous birds species - simply did not have multiple decades to make their reproductive mark. Instead the stakes were high to make their mark when they could and for many species had much less than a decade of good potential mating years. For large and combative males this opportunity may have been substantially less.

Abelisaurids - and by extension perhaps spinosaurids, unenlagines and several other rugose skulled dinosaurs - may have dispensed with scales on the head partially or completely and sported a tight fitting shellac of keratinized skin. Convergent in design with crocodilians and wood storks such a design would offer durability and protection while potentially allowing sensitivity for animals that explored their world largely through their heads.

Much to the consternation of both scale loyalists and feather nazis the more basal we go in many dinosaurian groups the more anarchic and varied might be the integumentary structures. In derived forms feathers or scales seem to predominate the ectoderm in various lineages. In maniraptorans it is feathers, while in ceratopsids, hadrosaurids, and abelisaurids it is scales. However in more basal forms of each group we might expect a veritable riot of weird, diverse, and unexpected integumentary coverings. This prediction is met by the rich and varied ectodermal topography of Kulindadromeus, the potential for both quills and diverse scales & scutes in Concavenator, quills and scales in many ornithischians, and the noted diversity of outgrowths in many basal mammal families today such as xenarthrans.

In short there is a lot to learn. Both scale loyalists and feather nazis have their role to play but I think we might be converging on a world where both sides are too dogmatic and that nekkid skin has a role to play in this as well. As well as keratinized skin. And funky jumbled up, mish-mashed piles of xenarthran like dinosaurs with piles of scutes, filaments, quills and other things galore. For those that like precise laws and set in stone didactic guidelines on how to depict dinosaurs - especially earlier more basal forms in which the developmental pathways were not yet established rigidily - probably not a pleasant time for you right now. And people of such mind sets will probably find greater and greater consternation at the sight of increasingly odd and bizarre dinosaur reconstructions. SUX2BU.

Ok, ok are you still with me? Have I done a good of enough job in triggering scale loyalists and feather nazis? Ok time to trigger myself a bit and it is the consistent critique I always get from some quarters regarding fleshy, hangy things or big lips jutting off the faces of theropods - especially toothy ones. triggrd

"But precious, such features would get snipped right off?!?"

I'm triggrd.
And this is my final word on this.

1) Don't underestimate how tough and flexible skin can be. Chewing on the wattles of a theropod could have been like chewing on a flat tire. You won't really hurt it and you might compromise your position allowing for a more devastating counter strike to the back of the neck, throat, or belly/cloaca.

I really like the thickness, elasticity, and well toughness in J.W. Kirby's depiction of SANTAnaraptor below... they look like tough chew toys that would stretch out and cause frustration to anything that bit into them.

SANTANAraptor credit J.W. Kirby

2) Critiques of soft tissue display/thermoregulatory structures on toothy theropods should familiarize themselves with the the "handicap" hypothesis of social-sexual signaling structures. In order for a signal to be true and just it must impart a cost on the signaler. That the signal actually imposes some maladaptive traits on the signaler is actually a good thing. Such signals represent the "conspicuous consumption" of the animal kingdom. They also should be expected in animals in which the stakes are high reproductively. And the stakes were definitely high reproductively in dinosaurs. They did not have several decades to make their mark in the Darwinian sweepstakes but only several years. In the largest and most combative large theropods how many years would you expect a large male to reign supreme? 2-3 years? Dinosaur sexual politics was a wild and wooly world, not for the feign of heart.

Also don't be carnivore biased. If you can accept fleshy, hangy wattles, dewlaps on ornithischians, herbivorous theropods, and sauropodomorphs but not on predatory theropods you are being biased. The giant beaks of ornithischians, ceratopsids, the powerful beak and jaws of an oviraptor, a macronarian are not to be trifled. These animals could easily rip your limbs from their sockets and could do the same to any fleshy display device. Should we eschew such devices on all dinosaurs then? - clearly not. In fact Edmontosaurus and Pelacinimimus prove that they had them!!

3) Male bull elephant seals and hooded seals engage in epic biting battle yet maintain fleshy, vulnerable facial outgrowths. Elephant seals form huge colonies... at least some maniraptorans did too. Male elephant seals have a severely limited time to get to the top of the social heap and mate. Theropods - indeed probably all dinosaurs - did not live as long as crocodiles so they were likely under the gun to achieve mating success at any cost. Male southern elephant seals are the largest mammalian carnivoran of all time. Now that is not stated enough in my opinion. The largest mammalian carnivoran of all time is not from the Eocene, Miocene, or early Pleistocene but living with us on earth right now. And it engages in absolutely vicious face biting prolonged skirmishes. And it has the size, strength, and tusks to do some real damage. Just like theropods did. Yet this animal - colony forming, combative, multi-ton largest carnivore ever - it has big, fleshy pendulous display structures right there on the tip of it's face. And you know that sometimes it gets ripped into, indeed it is a bit of a liability in fights but to be a true signal, to evince handicap, to resonate intimidating sounds and visually stimulate opponents - evolution has eclipsed these liabilities and favored large, fleshy pendulous things on the face of an animal that should rightly have them yanked right off. But it has them anyways and most likely did toothy theropods.

And finally - inb4 criticism - highly perceptive and critical reader will have detected what I have done here. By highlighting the need of scale loyalists to look at insulatory and thermal potential in scaley dinosaurs and by suggesting nekkid skin as a kinda proxy for a formerly fully feathered condition, I have deftly played both sides of the aisle and bolstered my claims of 1) S.I.G.I.L. i.e. insulation without feathers and 2) a fairly common evolution of facial nekkid skin and nekkid skin structures in feathered theropods. Duane you self serving bastard!!

34 comments:

Hi. One very quick comment -- virtually none of those things said about the integument of Pelecanimimus back in the day are actually true - I mean, they're not visible or not preserved, or have been misinterpreted. This doesn't affect your point, but it's worth noting.

Top post Duane - really enjoyed it.I have to admire the fact that you are one of the first guys to actually call out the Feather Nazis but I feel it will fall on deaf ears because they really are fanatical and will seldom enter into a constructive conversation - they are always right don't you know?

People have to understand that the truth is that no skin covering is mutually exclusive. I have also championed secondary featherless as well and I think this hypothesis has some merit. Yutyrannus is often cited as a potential pointer for the likelihood of feathers in large tyrannosaurids but this is actually hard to quantify since there is a significant temporal gap of around 40 million years between the two and not a lot of fossil evidence to help us out.So why not secondary featherless?

Phylogenetically, however, all the evidence suggests that tyrannosaurids were likely to sport some form of partial covering and you have to concede that a lot of phylogenetic predictions (but not all) have really come to the fore in recent years and have proven correct.

Still hard evidence remains elusive except for the fact that small scales are definitely known for tyrannosaurids and we should remember the Tarbosaurus mummy from Mongolia which was extensively covered in scales. However, as we all should, I remain open minded and will yield to the fossil evidence as it becomes apparent.

One last word for you. I cannot give much away but there is research going on right now alluding to something you mention in your post which will have you smiling all day long and shows good foresight on your part.

Very good post Duane. This might be one of my favorites. So much information about how utterly diverse Dinosaurs are by giving various lines of facts, reasoning and examples. The only thing even remotely wrong with it was the odd ball beginning comparing to Lord of the Rings and poor old Smeagol...that probably strayed away a few readers to be honest...but everything else was utterly spot on and incredibly intriguing and helpful.

And I too have been trying to usher in a guild of mixed bald, caruncled, scaley, feathere, etc. Archosaurs for a while now. Peple usually shrug me off but hey, with any ideological spectrum, the extremes of both are equally as ignorant. Too long have I seen completely scaley Maniraptorans AND competely dapper, ground hawk Maniraptorans. It's comical on both extremes of the spectrum really. Why do people insist on only giving fluffy Hawk and song bird feathers to any Terrestrial feathered Dinosaur? When was the last time a Ratite had soft fluffy feathers? The only exception for Ratites would be Tinamous...and it just so happens to be volant...I honestly get a lot of hate at time for giving anything remotely bald or simply "not feathered" to anything Maniraptoran. I don't draw much anymore, but Tyrannosaurs for instance I always give them a big, gnarly head of Caruncles, long shaggy Emu or Cassowary feathers on the body, a bald or scaley underbelly, scaley feet with thick pads and a tail that is to be variable. Sometimes scaley, sometimes bald, sometimes shaggy feathered and on occasion even quills. That's my average medium to large sized honestly. I love to give my Sauropods dewlaps as well since they'd be emitting so much body heat.

Also, didn't Andrea Cau make a post ages ago about how the "scaley" skin of Carnotaurus on the body wasn't actually armoured and scaley but rather flaky like a bald bird? Try to find his post about it, because if so then it means the body was likely feathered, or at the very least naked and wrinkly ala Elephant.

Yes, I did write a post on the actual evidence on the skin of Carnotaurus, based on the scientific description of the specimen (Bonaparte et al. 1990). It is funny how often people depict dinosaurs based on other depictions and not on actual scientific studies...

Duane, nice post. I think you will find this old picture I used in my blog more than 7 years ago very interesting: http://1.bp.blogspot.com/_6fVePgcYTJc/ScIzAWdf9VI/AAAAAAAABys/iBlmreKVk3Q/s1600/Archosauria+tegumentary+evolution2.jpgIn short, we must think in term of tegumentary gradients, not just two competing alternative conditions ("the reptilian vs the avian").

@Andrea Cau what is the conclusion for the body of Carnotaurus then? I saw how flaky it was and assumed a very bird like appearance. But weren't there also pockets where osteoderms would be connected every so often?

@mark wildman Thanks. Always happy to kill nazis lol. Again, tongue in cheek, and there are a lot of self described "feather nazis" that I like and respect. Its just getting too hemmed in to either end of the spectrum that becomes defeating. And you could say that many "feather nazis" are specifically intolerant of just JP stylized raptors however sometimes there is thought drift...

@khalil beiting. Thanks. But I particularly like the Lord of the Rings meta-analogy. Let it simmer a bit. I thought you would too being the meme king!!

@Andrea Cau Thanks and I actually remember that old picture, I do like how the ornithischian side, as it gets more derived moves away from filaments end of the spectrum while theropods do the opposite.

Active Wikipedia editor here. Have these observations regarding about preserved integument (or lack thereof, I should say....) in Pelecanimimus been published in the literature? Otherwise, it falls a little bit under the umbrella of original research, which is against Wiki policy....

I may be completely oblivious to it, but I never heard a "feathernazi" calling himself seriously. They always were a form of ironic humor to me, and I never heard of Feather millitantism. On the other hand, I did read many biased people defending scales for the sake of scales, and disregarding actual evidence. So I think this dichotomy is a fals one and is promoting a war-like mentality that is already too present in our world, mostly thanks to social media. But again, apart from your writing style that I subjectively dislike, you raise very good arguments and provoking thoughts, even though I was already sold on the weirdness of Abelisaurids and their facial integuments. And the concept of "soft dinosaur revolution" was already coined in the late 2000's, that we can link to the rise in "weirdness" you are talking about. I also agree that fleshy wattles are not impossible for animals with a violent lifestyle. To sum up, still don't like your style but respect your work.

that is what it says on the wikipedia page. there is a citation to a paper by D. E. G. Briggs, P. R. Wilby, B. Pérez Pérez-Moreno, J. L. Sanz, M. Fregenal-Martinez, (1997). "The mineralization of dinosaur soft tissue in the Lower Cretaceous of Las Hoyas, Spain." Journal of the Geological Society London, 154: 587-588.

According to Darren Naish, above comments, the interpretations (that I got from wikipedia I actually don't have the original paper) are ambiguous so I will take his word. I would like to more about what is going on there with Pelecanimimus myself...

One of my personal pet peeves is how much people rail on about Jurassic World needing to update their dinosaurs to better reflect modern scientific data out of a moral obligation to science. The only reason to think this is because you don't understand how copyright laws work and how tricky and weird it is to have to merchandise a franchise based on factual real animals.

The skin angle is wonderfully enlightening (especially that Croc thing. Very cool stuff!) The Abelisaurids are a favourite of mine and it's cool to see you finally touch on them.

Is it possible that there were "Southern" variations of T.Rex that tended towards nudity to accommodate heat with more appropriately woolly northern ones? The hubris of some people (often not formally educated in Paleontology BUTTHATSNONEOFMYBUSINESS) to think that the anarchic world of biology and animals could so conveniently fit into their simple categorizations and dichotomies. Did you learn nothing from Jurassic Park?

Great post! I think a lot of former "feather nazis " have been coming around to this way of thinking, but publication lags might make it look like the pendulum is still swinging towards "the enfluffening". I have dozens of illustrations of weird skin, combo skin, flashy fleshy patches, etc. in a variety of archosaur lineages, but they're earmarked for books and it'll be a few years ;)

@Duane Nash oh don't get me wrong, I love myself Lord of the Rings, The Hobbit or any other fantasy novel (and don't forget the memes!!!). It's just that many people may be off set when expecting a post about Archosaur integument to have a playful jab at many ala a reference to a random franchise! Amazing post regardless and I'm very excited for more, especially regarding integument.

IIRC, feathers are useful for soaking up a lot of heat and blocking external heat (though this doesn't address internal heat). While some birds have lost feathers for thermoregulation (discounting the wattles and display features of galliforms), most of these tend to be soaring flyers, and there it's a mixed bag

Khalil, there is not evidence of osteoderms in Carnotaurus.Osteoderms are present just along the dorsal midline in Ceratosaurus and eventually Eoabelisaurus. Given that recent analyses suggest Eoabelisaurus as a ceratosaurid, not an abelisauroid, the dorsal string of osteoderms may be restricted just to ceratosaurids.

We need more fossils and more careful description of the fossils before stating something robust on the tegument of most dinosaurs.

I agree with William and Matthew by saying that the so-called 'feather nazis' are more satirical than fanatics (or, at least, the ones I know of) and things are already changing for some time now - but still, doesn't affect your points!

The importance of skin structures and dewlaps is incredibly underrated and even if I'm not personally sure where the boundary between outrageously-weird and rightfully-weird ends (like we already discussed in a previous post of yours), we still need to remind how widespread these kind of structures are and how the dichotomy between Scales v. Feathers v. Naked skin shouldn't be a battle from a fantasy novel like you've said.

Thanks for all the comments and kind words. I'm not entirely comfortable with all this agreement and consensus building my instinct towards rebellion causing my pains… Yeah I mean the feather nazi thing is satirical and I get it. I think for those of us that remember when the Chinese revelations came about it was so exciting and opened up so many vistas of further exploration. We may have seen that wave reach its apogee and start to recede a bit so it feels a bit crestfallen. However as I made the point several times in the post scales/scutes/nekkid skin/keratinized skin are also interesting and may hold fruitful promise in unique and intriguing thermoregulatory/display benefit. Also things like the "bearded" filaments we see on turkeys which I learned our keratin outgrowths and not from feathers still offer a fruitful promise for that fabled somewhat filament covered sauropod… not there for insulation purpose but display!!

created an insulatory vacuum sealed layer when blood was withdrawn internally

How would that have worked? There is no way to withdraw blood and leave a vacuum behind. Empty blood vessels just collapse under the pressure of the air or water outside the organism.

(That's not an argument against naked skin here and there, of course.)

Consider xenarthrans - the strange and alluring mammals that include the extinct glyptodonts and giant grounds sloths, as well as armadillos, sloths, anteaters, probably some other weird ones too I am forgetting. I don't like to use the term "living fossil" but I think it fair to call these mammals the least derived or most basal among extant mammals. The most basal mammals but also the most diverse in terms of integumentary structures.

What, if anything, do you mean by "basal"? Xenarthrans are placentals, and they're the only synapsids with osteoderms. If the diadectomorphs are theropsids, well, they don't have osteoderms either. The osteoderms of xenarthrans are not a plesiomorphy, they're an apomorphy, and so are the epidermal scales that sometimes cover them.

Likewise, the scales of pangolins are apomorphic. Pangolins and carnivorans are sister-groups nested high up in Laurasiatheria.

==========

More (probably) later. For now, could you please work on your distinction of unstressed vowels in scientific terms? Aucasaurus, Pelecanimimus, pachycephalosaurids, seriema... an "Aucusaurus" with "oo" churns my stomach.

So what you think might have and probably was happening was that many non-avians had feathers, lost them, and the nekked skinned areas of their bodies would be something along the lines of elephants, bare skin but thick and sturdy. It was a little odd at first but it's growing on me. Happy New Year btw.

@David the idea I am referring to is S.I.G.I.L. which I linked to in the post. Follow the link to see further explanation of the idea. Maybe I was not clear enough but the idea is to have a keratinized, scaly outer layer for this to work. If you have a keratinized "scaly" outer integumentary structure and withdraw blood away from the epidermis won't those structures maintain their structural fidelity? Crocs can pump into and withdraw blood from their dorsal scutes, the scutes don't collapse when the blood leaves. If you have a hollowed out area under the scale it is now an insulation layer.

On xenarthran integumentary structures: Good points. Perhaps not the best examples then. However the idea that the more basal we go in many of these groups the more varied the integumentary coats might be is still a good idea worth exploring. Kulindadromeus seems to suggest this!!

On grammar errors: Uggh probably lots of errors. Probably not gonna change. Probably doing it a bit on purpose to elicit a response. Definitely working. Just a blog.

@Dromo Sapien Happy New Years to you too. Basically, but I don't want even the ideas I suggest to veer into dogmatism. I don't want people to make it a rule that naked headed maniraptorans are the expected look anymore than fully feathered headed maniraptorans are the rule either. No absolutes. There are some trends but even with these trends there are exceptions.

I myself may not be as knowledgeable in biology and paleontology as many of the readers here but I just wanted to say that this post was a very informative read and I find the idea of dinosaurs possessing a variety of integument types very exciting. Your posts always make for fine reads and you bring up plenty of "outside-the-box" ideas that I really don't see in many other paleontology circles. I'll make sure to share this and several other posts with friends and family online. It may bring back interest in some of them, who knows.

Regarding bird skin and dinosaur skin, your articles have really opened up so many possibilities. I can't figure out how to post images on this platform, but please Google Image Search for "Dong Tao Chicken".

Overall good post. You've put into writing what I've been saying for years. In fact your sketches at the end look very much like how I've pretty much always been drawing tyrannosaurids, albeit with more crazy cranial ornamentation.

Though I do have to agree with Matt and some others above that you're essentially attacking a strawman or a group of people that don't actually exist, but it still serves its function of getting the greater point across, so whatever.

We just need to get Emily Willoughby to paint an ugly, caruncled maniraptoran one of these days; if there are indeed any militant feather nazis out there, that should get them to come around. (She was the one who first converted me from scale loyalism!)

Somebody in the comments mentioned the phrase "Soft Dinosaur Revolution" (my eyes are glazing over as try to re-skim the comments to remember who): I think the term "Dinosaur Enlightenment" has more of a ring to it.

Emily might be a tough nut to crack on this one she seems pretty endeared to feathers but I would happily be proven wrong!!

Soft tissue revolution, soft dinosaur revolution, have all been advanced. This "Dinosaur Enlightenment" has a good connotation. What I particularly like about now is that it is not dominated by one or two thinkers as opposed to the earlier eras; there is a rich and fertile breeding ground for new ideas and depictions via the internet; and open access and the openness of many primary researchers has created a very interesting and somewhat novel collaboration between all strata of dinosaur and paleontology enthusiasts, lay people, amateurs, professionals. Of course there are the David Peters and such, you have to take the good with the bad, but I do think that dinosaur science - perhaps more than many other sciences - is showcasing how all these disparate levels and "strata" of entities can work together to further things; advance new hypotheses, ask questions, debate etc etc. It is a bit sloppy and haphazard but I feel we are on the cusp of imagining new ways that science can work, be communicated, and really a way people at many levels can engage actively in it.

I just have to note one thing: "strawman" does not remotely describe the individuals I've seen in full comment-chain froth on sites such as Deviantart, belaboring other artists-even artists whose stated intent was to experiment with dinosaur-styled fictional monsters, as opposed to accurate paleo depictions. Any given trend with a little traction will have its dogmatic extremists, and I'd say our host has been underplaying their Greater Internet F@#$@%$ Theory(C) aggro level if anything. Granted, some of the commenters above may have been too busy actually contributing to the field to notice, but trust me: there is an opposite extreme to the sort of blinkered perspective which curates Creationist museums.

@Carliro Good points as one commentator above has already pointed out. Indeed I am guilty of too hastily putting forth such an ill-equipped comparison. However I do think that as we go more basal into many of these dinosaur groups we will be confounded with more diversity and unexpected structures and combinations there of.

@SeanH. Indeed. There are multiple factions and I like to use the term "strata" of proponents on multiple sides to various degrees. I do mean no ill-will towards any of them. If anything my take home message to all is to be a little less dogmatic on all angles.

It's funny, because the thing that rustles me in this post isn't the feathers or lack thereof, but the discussion on them being brightly colored. I've even made a comment on it here before, lol.

To avoid making the same comment twice, I'll just say I think it's a bit too easy to assign vivid colors and modern bird display structures on non-avian dinosaurs when they had evolutionary stresses placed on them that modern avians haven't. Not saying they weren't there, just that people are too cavalier about it. Based on the prehistoric color patterns we have assembled based on scientific research, ecological niche seems to be a much bigger indicator of coloration possibilities than what modern relatives are currently sporting. As things are now, a lot of people are depicting dinos with a niche fitting a jaguar/lion/grizzly with a distinctly turkey/cassowary/peacock color scheme.

Moving on though, I think ultimately though that dinosaur feather patterns will prove much more varied than modern birds.

Tails would provide a significant amount of surface area for releasing excess heat that birds don't have access to. I don't think it's a coincidence that we have several feather patterns that lack feathers on or under the tail. The tail also at least in many cases circumvents the ratite use of wings in maintaining balance while running. They also didn't need them for flight (obviously). Any relatively small-armed theropod wouldn't need feathers on their arms, and dropping from those two locations may be more efficient than dropping feathers on the head and neck.

@Mr. Strongman Who is talking about brightly colored feathers? The post was mainly about naked skin (especially on head and neck) as a third option usually ignored by arguments over feathers vs scales. I don't know man are you commenting on the right post? This piece is not talking about bright coloration and neither are any of the above comments so…. I'll be talking about bright colors/display in theropods more in the future but I really doubt any need to infer the drab colorations of mammalian predators

I wasn't speaking specifically to feathers, but all display structures including crests, bills, horns, skin, etc. I probably could have been clearer on that, but you absolutely do have a section on display structures (including talking about hornbills, ground hornbills etc. as the closest matches to non-avians).

But, even if those are the closest matches, those animals in most cases aren't direct matches, and in all cases modern birds are either capable of flight or secondarily flightless. That means that the thoughts about things like thin bird skin I feel can be a trap because they would have had the pressure to drop weight wherever possible while non-avians that never became capable of flight never would have had to deal with such an issue -- particularly the larger grazers and browsers.

Flight also would have done two things to skew modern birds toward bright color displays (whether they are skin, scales, feathers, caruncles etc.)-- 1. birds developed as a clade very keen eyesight, skewing any sort of intra-species communication toward using visual cues instead of other senses and 2. gave most of them a defense mechanism which would have allowed them better defense against predators and reducing the need to have more subdued/camouflage colorations.

Yes, but we do have direct bony evidence of display structures in theropods (e.g. Dilophosaurus, Monolophosaurus etc etc). All I'm saying is that there were probably loads of skin (and feather) derived display structures too in addition to the ones we know of for sure.

Are you suggesting that, with regards to color, that even for the display structures we know (from bony evidence not just speculation) for certain that they were not brightly colored but more subdued in coloration? Is that what you are suggesting? Just for clarity because I do get this suggestion from others too... and following this you seem to suggest that "birds developed as a clade very keen eyesight" which, correct me if I'm wrong, sort of implies dinosaurs and non-avian theropods did not have color vision?

I mean just for clarity are you suggesting a very drab dinosaur display culture? Sort of more like the tans, greys, and browns of large mammals? With the exception of say certain color vision capable primates...